Domestication and exposure to human social stimuli are not sufficient to trigger attachment to humans: a companion pig-dog comparative study.

Dogs exhibit human-analogue attachment to their owners, with similar function and mechanisms to that of infant-mother bond, but its origin is unclear. Comparative studies on socialised wolves and dogs emphasise genetic influence in dogs' preparedness for attachment to humans. We aimed to reveal if this genetic effect stems from general domestication or artificial selection that increased dogs' dependence on humans. We assessed and compared behavioural patterns of young companion pigs and dogs using a Strange Situation Test. Dogs but not pigs exhibited distinct behaviours towards their owner and a stranger along attachment-specific variables, so only dogs' relevant behaviours fulfilled attachment criteria. From the observed behaviours, three factors were formed: Attachment (to the owner), Anxiety (in a strange situation), and Acceptance (of a stranger). Results indicate (1) higher Attachment scores in dogs than pigs, (2) greater Acceptance scores in pigs, (3) positive correlation of Attachment and Anxiety in both, (4) similar time tendency of pigs' Attachment and Acceptance scores. These suggest that in pigs, domestication and early exposure to human social stimuli did not trigger attachment to humans. Thus, along with species predispositions, the unique dog-owner attachment can be facilitated by artificial selection that increased dogs' dependence on humans.

Out-of-reach rewards elicit human-oriented referential communicative behaviours in family dogs but not in family pigs.

Human-oriented referential communication has been evidenced not only in domestic but also in some wild species, however, the importance of domestication-unrelated species' characteristics in the emergence of this capacity remains largely unexplored. One shared property of all species reported to exhibit referential communication is the efficient use of visual social signals. To assess the potential role of species-specific characteristics in the emergence of human-oriented referential communication, we compared similarly socialised companion animals from two domestic species: dogs, which rely heavily on conspecific visual social signals; and pigs, which do not. We used an out-of-reach reward paradigm with three conditions: both human and reward present, only human present, only reward present. Both species exhibited certain behaviours (e.g. orientation towards the human, orientation alternation between the human and the reward) more often in the human's presence. However, only dogs exhibited those behaviours more often in the simultaneous presence of the human and the reward. These results suggest similar readiness in dogs and pigs to attend to humans but also that pigs, unlike dogs, do not initiate referential communication with humans. The ability to referentially communicate with humans may not emerge in mammals, even if domesticated companion animals, that lack certain species characteristics, such as efficient intraspecific visual communication.

The acoustic bases of human voice identity processing in dogs.

Speech carries identity-diagnostic acoustic cues that help individuals recognize each other during vocal-social interactions. In humans, fundamental frequency, formant dispersion and harmonics-to-noise ratio serve as characteristics along which speakers can be reliably separated. The ability to infer a speaker's identity is also adaptive for members of other species (like companion animals) for whom humans (as owners) are relevant. The acoustic bases of speaker recognition in non-humans are unknown. Here, we tested whether dogs can recognize their owner's voice and whether they rely on the same acoustic parameters for such recognition as humans use to discriminate speakers. Stimuli were pre-recorded sentences spoken by the owner and control persons, played through loudspeakers placed behind two non-transparent screens (with each screen hiding a person). We investigated the association between acoustic distance of speakers (examined along several dimensions relevant in intraspecific voice identification) and dogs' behavior. Dogs chose their owner's voice more often than that of control persons', suggesting that they can identify it. Choosing success and time spent looking in the direction of the owner's voice were positively associated, showing that looking time is an index of the ease of choice. Acoustic distance of speakers in mean fundamental frequency and jitter were positively associated with looking time, indicating that the shorter the acoustic distance between speakers with regard to these parameters, the harder the decision. So, dogs use these cues to discriminate their owner's voice from unfamiliar voices. These findings reveal that dogs use some but probably not all acoustic parameters that humans use to identify speakers. Although dogs can detect fine changes in speech, their perceptual system may not be fully attuned to identity-diagnostic cues in the human voice.

Occurrences of non-linear phenomena and vocal harshness in dog whines as indicators of stress and ageing.

During social interactions, acoustic parameters of tetrapods' vocalisations reflect the emotional state of the caller. Higher levels of spectral noise and the occurrence of irregularities (non-linear phenomena NLP) might be negative arousal indicators in alarm calls, although less is known about other distress vocalisations. Family dogs experience different levels of stress during separation from their owner and may vocalise extensively. Analysing their whines can provide evidence for the relationship between arousal and NLP. We recorded 167 family dogs' separation behaviour including vocalisations, assessed their stress level based on behaviour and tested how these, their individual features, and owner reported separation-related problems (SRP) relate to their whines' (N = 4086) spectral noise and NLP. Dogs with SRP produced NLP whines more likely. More active dogs and dogs that tried to escape produced noisier whines. Older dogs' whines were more harmonic than younger ones', but they also showed a higher NLP ratio. Our results show that vocal harshness and NLP are associated with arousal in contact calls, and thus might function as stress indicators. The higher occurrence of NLP in older dogs irrespective to separation stress suggests loss in precise neural control of the larynx, and hence can be a potential ageing indicator.

Who turns to the human? Companion pigs' and dogs' behaviour in the unsolvable task paradigm.

When facing an unsolvable problem, dogs exhibit spontaneous human-oriented behaviours (e.g. looking at the human partner, gaze alternations between the human and the target) sooner and for longer than domestic cats and hand-raised wolves. These behaviours have been interpreted as interspecific communicative acts aimed to initiate interaction. Here, we compare the emergence of human-oriented behaviours (e.g. orientation towards humans, orientation alternations, vocalizations) in similarly raised family dogs and miniature pigs utilising an unsolvable task paradigm which consists of Baseline (no task), Solvable and Unsolvable phases. Relative to the Baseline phase in which both species showed human-oriented behaviours to a similar extent, during the Unsolvable phase dogs showed more and pigs showed less such behaviours. Species-predispositions in communicative behaviour may explain why dogs have a higher inclination than pigs to initiate interspecific interactions with humans in problem-solving contexts.

Human proximity seeking in family pigs and dogs.

Family dogs (Canis familiaris) seek human contact from an early age, can discriminate and prefer their caregivers over other humans. To investigate if being kept as a family animal is sufficient to induce similar early human proximity-seeking in another domestic mammal, here we directly compared such behaviours in dogs and similarly raised domestic pigs (Sus scrofa domesticus). We used a preference test to measure proximity-seeking behaviours of 4-month-old family pigs and dogs in the presence of their caregiver and either a stranger or a familiar object, in a novel environment. We found that both pigs and dogs preferred their caregivers over the familiar object but not over the stranger. However, when the caregiver and the stranger were present, only dogs showed an overall preference for human proximity, and pigs spent more time away from both humans. These results suggest that both dogs and pigs seek the proximity of their caregiver, but there is a difference in how each species generalizes their experience to other humans. Species-specific predispositions, including that dogs have a longer socialization period and that humans are more salient as a social stimulus for them, may be important for the early development of an overall preference for humans.

Comparing interspecific socio-communicative skills of socialized juvenile dogs and miniature pigs.

Previous research proves dogs' outstanding success in socio-communicative interactions with humans; however, little is known about other domestic species' interspecific skills when kept as companion animals. Our aim was to assess highly socialized young miniature pigs' spontaneous reactions in interactions with humans in direct comparison with that of young family dogs. All subjects experienced similar amount of socialization in human families. In Study 1, we investigated the appearance of human-oriented behaviours without the presence of food (Control condition) when a previously provided food reward was withheld (Food condition). In Study 2, we measured responsiveness to two types of the distal pointing gesture (dynamic sustained and momentary) in a two-way object choice test. In the Control condition of Study 1, the duration of pigs' and dogs' orientation towards and their frequency of touching the human's body was similar. In the Food condition, these behaviours and orienting to the human's face were intensified in both species. However, pigs exhibited face-orientation to an overall lesser extent and almost exclusively in the Food condition. In Study 2, only dogs relied spontaneously on the distal dynamic-sustained pointing gesture, while all pigs developed side bias. The results suggest that individual familiarization to a human environment enables the spontaneous appearance of similar socio-communicative behaviours in dogs and pigs, however, species predispositions might cause differences in the display of specific signals as well as in the success of spontaneously responding to certain types of the human pointing gestures.

Investigating jealous behaviour in dogs.

The function of jealous behaviour is to facilitate the maintenance of an important social relationship that is threatened by a third-party, a rival individual. Although jealous behaviour has an important function in gregarious species, it has been investigated almost exclusively in humans. Based on functional similarity between dog-owner and mother-infant attachments, we hypothesised that jealous behaviour can be evoked in dogs, similarly to children. In our study owners focused their attention solely on the test partner, while they ignored their dog. We deployed familiar and unfamiliar dogs as social test partners, and familiar and unfamiliar objects as non-social test partners; all subjects encountered all test partners. Dogs showed more jealous behaviour, i.e. owner-oriented behaviour and attempts to separate the owner and test partner in case of social compared to non-social test partners. Results suggest that jealous behaviour emerges in dogs, and it is functionally similar to that in children observed in similar situations. Alternative explanations like territoriality, dominance rank can be excluded.